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WO1996039807A1 - Procede de developpement de plantes infectees par des endophytes - Google Patents

Procede de developpement de plantes infectees par des endophytes Download PDF

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Publication number
WO1996039807A1
WO1996039807A1 PCT/US1996/009278 US9609278W WO9639807A1 WO 1996039807 A1 WO1996039807 A1 WO 1996039807A1 US 9609278 W US9609278 W US 9609278W WO 9639807 A1 WO9639807 A1 WO 9639807A1
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WIPO (PCT)
Prior art keywords
endophyte
bentgrass
turf grass
cultivar
grass
Prior art date
Application number
PCT/US1996/009278
Other languages
English (en)
Inventor
A. Douglas Brede
Suichang Sun
Original Assignee
Jacklin, Donald, W.
Jacklin, Doyle, W.
Jacklin, Duane, A.
Priority date (The priority date is an assumption and is not a legal conclusion. Google has not performed a legal analysis and makes no representation as to the accuracy of the date listed.)
Filing date
Publication date
Application filed by Jacklin, Donald, W., Jacklin, Doyle, W., Jacklin, Duane, A. filed Critical Jacklin, Donald, W.
Priority to AT96918248T priority Critical patent/ATE204121T1/de
Priority to JP9501633A priority patent/JPH11507216A/ja
Priority to EP96918248A priority patent/EP0836378B1/fr
Priority to DE69614548T priority patent/DE69614548T2/de
Priority to DK96918248T priority patent/DK0836378T3/da
Priority to AU60950/96A priority patent/AU696959B2/en
Publication of WO1996039807A1 publication Critical patent/WO1996039807A1/fr

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Classifications

    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
    • A01H4/00—Plant reproduction by tissue culture techniques ; Tissue culture techniques therefor
    • A01H4/005—Methods for micropropagation; Vegetative plant propagation using cell or tissue culture techniques
    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
    • A01H5/00—Angiosperms, i.e. flowering plants, characterised by their plant parts; Angiosperms characterised otherwise than by their botanic taxonomy
    • A01H5/12—Leaves
    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
    • A01H6/00—Angiosperms, i.e. flowering plants, characterised by their botanic taxonomy
    • A01H6/46—Gramineae or Poaceae, e.g. ryegrass, rice, wheat or maize
    • A01H6/466—Poa, e.g. bluegrass
    • Y—GENERAL TAGGING OF NEW TECHNOLOGICAL DEVELOPMENTS; GENERAL TAGGING OF CROSS-SECTIONAL TECHNOLOGIES SPANNING OVER SEVERAL SECTIONS OF THE IPC; TECHNICAL SUBJECTS COVERED BY FORMER USPC CROSS-REFERENCE ART COLLECTIONS [XRACs] AND DIGESTS
    • Y10—TECHNICAL SUBJECTS COVERED BY FORMER USPC
    • Y10S—TECHNICAL SUBJECTS COVERED BY FORMER USPC CROSS-REFERENCE ART COLLECTIONS [XRACs] AND DIGESTS
    • Y10S47/00—Plant husbandry
    • Y10S47/01—Methods of plant-breeding and including chromosome multiplication

Definitions

  • This invention relates to new cultivars of bentgrass and Kentucky bluegrass which are stably infected with performance-enhancing endophytes and to methods of developing these cultivars.
  • the cultivars provide new turf grass varieties having increased resistance to damage by insect pests and certain fungal infections and increased tolerance to environmental stress such as summer heat and drought conditions.
  • BACKGROUND Endophytes of the genus Acre/ ⁇ oniiun are beneficial fungi which live within their plant hosts. They exist within the plant for all or nearly all of their life cycle. Endophytes are found within the leaves, stems and flowering parts of healthy plants, generally lack a sexual stage in their development, and have limited, if any, pathogenic effects on their plant hosts. Plant biologists have been aware of the presence of endophytes in many natural grasses for more than 100 years. However, they have only recently begun to appreciate the role that these fungi play in enhancing the performance of certain grasses, including those that are useful for conservation of natural landscapes and for turf on lawns.
  • U.S. Patent No. 4,940,834 discloses a method for developing a new variety of perennial ryegrass in which a selected, naturally occurring maternal seed parent, which hosts an endophytic fungus capable of enhancing its host plant's characteristics, is crossed with a paternal pollen parent plant of the same or different variety as the maternal parent. Progeny from that cross, which display the desired endophyte associated characteristics, are selected for propagation by any of the methods of plant propagation presently used by those of skill in the art of plant breeding. These methods include paired hybridization, polycross hybridization and recurrent selection.
  • endophytes do not naturally occur in certain plant species, particularly in popular turf grasses such as the bentgrasses of the genus Agrostis and Kentucky bluegrasses of the genus Poa .
  • a recent field survey by White et al. found endophytes in only four species of the thirteen species of
  • Plant biologists have attempted to develop methods for directly incorporating endophytes into grass cultivars, so that the performance-enhancing characteristics resulting from natural endophyte infection can be introduced into desirable, elite turf grasses such as bentgrass and Kentucky bluegrass.
  • These approaches have involved attempts to inoculate seedlings with endophytes directly, as well as, attempts to introduce endophytes into callus grown in tissue culture, into somatic embryos isolated from callus tissue and into plantlets developed in tissue culture from meristems.
  • endophytes into callus grown in tissue culture, into somatic embryos isolated from callus tissue and into plantlets developed in tissue culture from meristems.
  • the present invention provides new cultivars of bentgrasses and Kentucky bluegrass that are stably infected with beneficial fungal endophytes. These cultivars have certain enhanced performance characteristics, including improved resistance to insects, resistance to diseases and tolerance to stress, which result in improved grass growth and survival under unfavorable environmental conditions such as summer heat and drought.
  • This invention further provides methods for infecting turf grasses with endophytes by direct inoculation.
  • the methods can be used to develop new endophyte-infected turf grasses having desirable characteristics, including improved resistance to insects, growth characteristics and tolerance to environmental stress.
  • novel endophyte-infected turf grass cultivars of this invention may be established initially by methods which involve directly incorporating endophytes into seedlings or other appropriate plant tissues of naturally occurring endophyte-free grasses. Once the plant tissues are infected, they are used for the development of novel turf grass cultivars having endophyte-enhanced performance characteristics. These novel cultivars are then used in more traditional plant-breeding procedures for producing further improved grass varieties.
  • Turf grasses which are not naturally infected by endophytes include those species known as bentgrass, of the genus Agrostis, particularly creeping bentgrass of the species Agrostis stolonifera, which is understood to be synonymous with Agrostis palustris and Kentucky bluegrass, of the genus Poa, particularly the species Poa pratensis, which is understood to be synonymous with Poa angustifolia, both of which are used in intensively managed turf surfaces such as lawns and golf course greens, tees and fairways.
  • novel endophyte infected turf grass may be developed by an elective crossing procedure.
  • Varieties of closely related grasses which are naturally infected with endophyte may selected in the wild as native, uncultivated grasses.
  • the species may include "wild" varieties of native bentgrasses. These species may further include "wild" Poa species such as Poa arida, Poa nervosa , Poa sterilis, Poa glauci folia, Poa cuspidata, Poa arctica, Poa ampla as well as other wild species of Poa .
  • the inventors of the present application have identified a number of such endophyte- infected species of Poa which are related to Kentucky bluegrass, Poa pratensi ⁇ , and which have a similar morphology and appearance. Surprisingly, it has now been found that endophytes can be introduced into Kentucky bluegrass by crossing with these selected wild-type species.
  • Endophyte-infected plants of wild-type Poa are brought into the greenhouse and propagated, then hybridized with cultivars of Kentucky bluegrass, the infected wild-type plants being used as the maternal parents. Endophyte-infected segregating progeny of this hybridization having dominate characteristics of Kentucky bluegrass are selected for further propagation.
  • Selected grass varieties can be infected with Acremonium endophytes as seedlings, as callus tissue, as plantlets derived from single meristems or as somatic embryos, using methods known to those of skill in the art of plant tissue culture. Methods of culturing plant tissues are well known to modern plant biologists. See for example, Johnson, Bush, and Siegel, "Infection of Tall Fescue with Acremonium coenophialum by Means of Callus Culture," Plant Disease 70. (1986) pp 380-382.
  • Plantlets for inoculation can be prepared from single meristems of mature bentgrass or Kentucky bluegrass plants using procedures analogous to those described for perennial ryegrass plants by O'Sullivan and Latch using grass meristems in tissue culture.
  • B. D. O'Sullivan and G. C. M. Latch "Infection of plantlets derived from ryegrass and tall fescue meristems, with Acremonium endophytes," in Proceedings of the Second International Symposium on Acremonium/ Grass Interactions, D.E. Hume, G.C.M. Latch and H.S. Easton, eds.
  • endophyte inoculations are carried out on grass seedlings.
  • seedlings is advantageous, because it allows the inoculation and screening of large numbers of individual plants, and therefore increases the chances of identifying endophyte-infected plants in inoculations where the efficiency of infection is low.
  • Infection of a single endophyte-free species or variety may require the inoculation of between 100 to 5000 seedlings. Preferably, between 1000 to 2000 seedlings are inoculated.
  • Specimens of native bentgrass ecotypes can be collected from plant populations in Europe and its neighboring islands, western Asia, and parts of the Middle East. Introduced populations can be found throughout all temperate regions of the world, including North America, where the grasses have escaped from cultivation.
  • the species can be identified by trained botanists or agriculturalists by the characteristic shape of the leaf blade, and the arrangement of the seed panicle at maturity. Alternatively, known, commercially available, endophyte-free turf grass varieties may be used in these methods.
  • Preferred, commercially available, endophyte-free turf grass varieties which may be used for developing the cultivars of the present invention include the creeping bentgrass varieties known as Putter, Southshore, Buffalo, SR1020, Pennlinks, Penncross, Penneagle, Emerald, Pro/cup, Crenshaw, Cato, 18th Green, and Cobra.
  • Preferred bluegrass varieties include those known as Adelphi, Amazon, Cheri, Classic, Eclipse, Freedom, Glade, Julia, Limousine, Nustar, Ram I, and Nublue.
  • Additional preferred bluegrass varieties include those known as Dawn, Destiny, Fylking, Huntsville, Liberty, Merion, Nassau, Nugget, Suffolk, Wabash, A-34, Able I, Classic, Monopoly, Shamrock, Merit, Barblue, Touchdown, Indigo, Blacksburg, Abbey, Coventry, Washington, Georgetown, Preakness, Livingston, P-104, P-105, 1757, Challenger, Midnight, Unique, Apex, Pennpro, SR2000, SR2100, Trenton, Opal, Gnome, Haga, and common.
  • Seeds to be used for the germination of seedlings are collected from any of the bentgrass or Kentucky bluegrass, particularly the varieties disclosed above. Seeds from commercial varieties of bentgrass and Kentucky bluegrass can be obtained directly from commercial sources.
  • Endophytes useful for infecting bentgrass seedlings are isolated from natural endophyte-infected grasses collected from nature in areas where the Agrostis genus is native and well adapted. Collecting endophytes from the epicenter of a species' origin provides the collector with plant specimens having increased genetic diversity. In theory, this occurs because the plants at the epicenter have had a longer time to evolve and are therefore more diverse.
  • endophyte-infected plants can also be expected to increase in number as one nears the species' epicenter, since the endophyte- host interaction is highly specific.
  • Compatible endophytes are identified only through methods of inoculation and subsequent microscopic examination. To date, there have been no tests developed by plant scientists to predict whether a given endophyte genotype will be compatible with a given grass genotype. Selection of a large number of possible endophyte candidates, and a large number of grass candidates greatly increases the possibility of a compatible match.
  • plant biologists Prior to the studies disclosed herein, plant biologists have been unable to find a satisfactory "match" between an isolated endophyte and a grass genotype for the bentgrass and Kentucky bluegrass species.
  • Collected native grasses and commercial grasses can be surveyed for endophytes by employing the culm examination technique. Culms, the jointed stem of the gra ⁇ s, are split longitudinally and the inner tissues stained with 0.5% rose bengal, then scraped onto a clean glass slide and examined microscopically. Similar procedures are used for screening inoculated seedling and plantlets grown from tissue culture to ascertain whether inoculation procedures have produced endophyte-infected specimens.
  • Endophytes may be isolated from infected tissues of collected grasses by procedures known in the art. For example, see J.F. White et al. , American Journal of Botany 79 (1992) at 473.
  • the surfaces of culms or leaf sheaths are preferably disinfected, e.g., by treating with a solution of 20% hypochlorite bleach solution, then rinsing in sterile water.
  • the disinfected plant tissues are then plated on an appropriate medium such as potato dextrose agar (PDA) .
  • PDA potato dextrose agar
  • endophyte strains are known to cause a plant disease known as choke, which effectively kills the developing grass seedhead, preventing seed production on those plants. These strains of endophyte are not suitable for use for commercial seed-propagated cultivars and should therefore be avoided when selecting strains of endophytes for inoculation. Likewise endophyte-infected wild-type grasses displaying the symptoms of choke should be avoided for use in selective hybridizations.
  • Plant tissues may be inoculated with endophytes by placing the endophyte into a direct wound or cut made in the plant tissue.
  • the cut may be made directly into the stem of the plant.
  • the inoculation is carried out by placing a small piece of an endophyte directly on or into the cut.
  • callus tissue may.be cut and the endophyte mycelium applied directly to the wound.
  • One preferred method which can be used on plant parts other than seed, utilizes the above- ground grass stolon, centering on the secondary shoot arising from the node as the tissue for carrying out the inoculations.
  • the inoculation with endophyte mycelium should be repeated from 1 up to 5 times, at approximately 7 to 10 day intervals after the previous inoculation has been done.
  • the repeated inoculation procedure is essential to successful incorporation of the endophyte, because it provides a greater • opportunity for the endophyte to penetrate the host before the host wound heals.
  • Inoculated plant tissues are allowed to heal, and the tissues are allowed to develop into plants which can be further cultivated by methods appropriate for the tissue type initially inoculated. If plantlets have been inoculated, they are grown in sterile culture for six to eight weeks before they are transferred to greenhouse and transplanted into soil.
  • callus tissue has been inoculated with endophyte, the tissue is transferred into auxin-free medium to allow the development of somatic embryos and plantlets.
  • Inoculated plants are removed from the sterile growth medium using forceps and transferred into (non- sterile) greenhouse potting mixture.
  • the plantlets are grown in potting mix for 4 to 6 weeks until they are of sufficient size that a piece of the stem material can be excised and checked under the microscope for the presence of the endophyte.
  • endophyte infection is maternally transmitted to progeny.
  • the present invention enables the preparation of a wide variety of infected cultivars by crossing with any other plant that is capable of breeding with an infected plant. Accordingly, the present invention provides a wide variety of endophyte-infected bentgrass and Kentucky bluegrass cultivars, whereas before this invention, infected plants of these species were unknown.
  • the invention is further illustrated by the following examples, which are not intended to be limiting.
  • Creeping bentgrass seeds were sterilized by a process known as double sterilization. The seeds were first washed with a solution of 50% hypochlorite bleach, then rinsed with distilled water three to four times. The seeds were drained and allowed to rest overnight. The entire sterilization process was repeated the next day so that any spores which germinated overnight were also destroyed.
  • the sterile seeds were transferred to a sterile Murashige and Skoog medium without plant hormones, and then allowed to grow until they were between 0.5 and 1.0 inches in length, which usually required between 7 to 10 days of growth.
  • Clones from the bentgrass variety Putter were inoculated with mycelium taken from a culture of endophyte identified as AS-7, which was obtained from Dr. James White of Auburn University.
  • the endophyte was originally isolated from plants growing in central England, the exact species of which has not been ascertained.
  • the seedlings were inoculated by making a small cut with a sterile scalpel into the stem of the plant, then laying an endophyte directly on top of, or into the cut. The process was repeated after 7 to 10 days when the wound began to heal. The process could be repeated additional times at 7 to 10 day intervals to increase the probability of successful inoculation.
  • the inoculation with endophyte mycelium is done immediately after each incision is made.
  • Inoculated seedlings were allowed to grow in petri dishes with sterile Murashige and Skoog medium for six to eight weeks after the inoculations. They were then moved to the greenhouse and transplanted into soil to allow the new plants to develop. Samples of tissue from the young plants were examined microscopically to determine if the endophyte had established itself in the cuttings after the plants have developed four or five tillers. After the endophyte has been confirmed within the plant, and reverified after 4 to 6 months of vegetative growth, then the infected plant is placed into a conventional plant breeding program. After inoculation, the plant line is usually too restricted or narrow in genetic base to constitute a complete and finished variety.
  • seed is harvested from a population of endophyte-containing plants in a breeder seed field. Each plant in the population has been checked under the microscope to confirm the presence of endophyte prior to anthesis. Any non-endophyte plants are discarded following pollination, and their seed is not harvested. In theory, this produces a variety with near 100% endophyte infection.
  • Bentgrass cultivars developed by the foregoing methods posses certain desirable performance characteristics, in addition to the presence of an endophyte. These cultivars have medium dark green genetic color, an upright leaf orientation, lack of steminess when grown as maintained turf, and good tolerance to close mowing, disease, drought, and stress. These cultivars also exhibit improved botanical insect resistance, in addition to the resistance provided by the endophyte.
  • Poa arctica were collected from native grass populations located in eastern Montana. The specimens were potted and brought into the greenhouse prior to seedhead expression, at which time their infection with endophyte was reverified. These species are related to Poa pratensis, Kentucky bluegrass, in that they have a rhizome habit and a similar morphology and appearance.
  • Hybrid progeny having Kentucky bluegrass morphology constituted from about 1% to about 30% of the total plants obtained from the selective hybridization procedure, depending on the mother plant used.

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Abstract

On sait que les endophytes utiles qui vivent dans certaines plantes permettent de lutter contre les insectes de façon biologique, économique et souhaitable. De nombreuses herbes présentes dans la nature hébergent des champignons endophytes symbiotiques. Cependant les endophytes utiles n'ont jamais été mis en évidence dans plusieurs variétés d'herbes à gazon, y compris des variétés d'agrostides et de pâturins des prés, deux herbes à gazon importantes sur le plan commercial et qui sont largement utilisées sur les terrains de golf et les gazons. L'invention concerne de nouvelles méthodes d'inoculation de tissus végétaux permettant le développement de variétés d'herbes à gazon, optimisées par des endophytes. Elle concerne également des variétés d'herbes à gazon produites au moyen de ces méthodes.
PCT/US1996/009278 1995-06-07 1996-06-05 Procede de developpement de plantes infectees par des endophytes WO1996039807A1 (fr)

Priority Applications (6)

Application Number Priority Date Filing Date Title
AT96918248T ATE204121T1 (de) 1995-06-07 1996-06-05 Verfahren zur entwicklung von endoptytischen- infizierten pflanzen
JP9501633A JPH11507216A (ja) 1995-06-07 1996-06-05 内生植物感染植物の発生方法
EP96918248A EP0836378B1 (fr) 1995-06-07 1996-06-05 Procede de developpement de plantes infectees par des endophytes
DE69614548T DE69614548T2 (de) 1995-06-07 1996-06-05 Verfahren zur entwicklung von endoptytischen-infizierten pflanzen
DK96918248T DK0836378T3 (da) 1995-06-07 1996-06-05 Fremgangsmåde til udvikling af endofytinficerede planter
AU60950/96A AU696959B2 (en) 1995-06-07 1996-06-05 Process for the development of endophyte-infected plants

Applications Claiming Priority (2)

Application Number Priority Date Filing Date Title
US08/483,897 US5723720A (en) 1995-06-07 1995-06-07 Process for the development of endophyte-infected plants
US08/483,897 1995-06-07

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US (1) US5723720A (fr)
EP (1) EP0836378B1 (fr)
JP (1) JPH11507216A (fr)
AT (1) ATE204121T1 (fr)
AU (1) AU696959B2 (fr)
CA (1) CA2223882A1 (fr)
DE (1) DE69614548T2 (fr)
DK (1) DK0836378T3 (fr)
ES (1) ES2160250T3 (fr)
PT (1) PT836378E (fr)
WO (1) WO1996039807A1 (fr)

Cited By (5)

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Publication number Priority date Publication date Assignee Title
EP0753259A3 (fr) * 1995-06-15 1997-05-28 Maekawa Seisakusho Kk Herbe et méthode d'introduction des champignons endophytiques dans l'herbe
US6072107A (en) * 1997-05-27 2000-06-06 New Zealand Pastoral Agriculture Research Institute Limited Ryegrass endophytes
US6111170A (en) * 1997-05-27 2000-08-29 New Zealand Pastoral Agriculture Research Institute Limited Tall fescue endophytes
US6815591B1 (en) 1999-04-16 2004-11-09 Advanta Seeds B.V. Enhancing endophyte in grass
CN108703068A (zh) * 2018-04-04 2018-10-26 广西壮族自治区农业科学院生物技术研究所 去除慈姑培养过程中内生菌的方法、培养方法以及应用

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CA2215849A1 (fr) * 1997-09-11 1999-03-11 Christophe Michot Nouveau solvant et composition electrolytique a haute conductivite ayant une stabilite a large plage
WO2000040075A1 (fr) 1999-01-05 2000-07-13 Advanta Seeds Pacific Inc. Endophyte et herbe infecte par ce champignon
WO2002013616A2 (fr) * 2000-08-15 2002-02-21 The Board Of Trustees Of The University Of Arkansas Endophytes non toxiques, plantes infectees avec et procedes d'infection de plantes
US20040172672A1 (en) * 2001-05-15 2004-09-02 Wilkinson Theresa C. Method for producing transgenic monocotyledonous plants
US7901927B1 (en) * 2005-08-09 2011-03-08 The United States Of America As Represented By The Secretary Of Agriculture Transfer and incorporation of heritable symbiotic fungi into non-host plants
US7642424B2 (en) * 2006-07-10 2010-01-05 Barenbrug Usa, Inc. Tall fescue endophyte E34
US8455395B2 (en) * 2006-10-24 2013-06-04 Irving Licensing, Inc. Endophyte enhanced seedlings with increased pest tolerance
US7892813B2 (en) * 2007-02-12 2011-02-22 The Samuel Roberts Noble Foundation Fungal endophytes of Elymus canadensis
CA2673696A1 (fr) * 2008-07-25 2010-01-25 The Samuel Roberts Noble Foundation, Inc. Amelioration de biomasse et de productivite dans des plants de gazon
US9469836B2 (en) 2011-01-28 2016-10-18 J.D. Irving, Limited Antifungal metabolites from fungal endophytes of Pinus strobus
CN109804913B (zh) * 2019-01-17 2021-09-28 山西大学 一种白羊草-内生真菌共生体的构建方法
CN113692963A (zh) * 2021-09-08 2021-11-26 上海水生环境工程有限公司 一种快速成型的藻-草共生水生草皮的培育方法及应用
CN114902914A (zh) * 2022-05-09 2022-08-16 兰州大学 一种野大麦-内生真菌共生体的构建方法

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Cited By (6)

* Cited by examiner, † Cited by third party
Publication number Priority date Publication date Assignee Title
EP0753259A3 (fr) * 1995-06-15 1997-05-28 Maekawa Seisakusho Kk Herbe et méthode d'introduction des champignons endophytiques dans l'herbe
US5880343A (en) * 1995-06-15 1999-03-09 Mayekawa Manufacturing Co., Ltd. Grass and method of introducing endophytic fungi into a grass
US6072107A (en) * 1997-05-27 2000-06-06 New Zealand Pastoral Agriculture Research Institute Limited Ryegrass endophytes
US6111170A (en) * 1997-05-27 2000-08-29 New Zealand Pastoral Agriculture Research Institute Limited Tall fescue endophytes
US6815591B1 (en) 1999-04-16 2004-11-09 Advanta Seeds B.V. Enhancing endophyte in grass
CN108703068A (zh) * 2018-04-04 2018-10-26 广西壮族自治区农业科学院生物技术研究所 去除慈姑培养过程中内生菌的方法、培养方法以及应用

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EP0836378A1 (fr) 1998-04-22
DE69614548D1 (de) 2001-09-20
CA2223882A1 (fr) 1996-12-19
DE69614548T2 (de) 2001-11-22
EP0836378B1 (fr) 2001-08-16
PT836378E (pt) 2001-12-28
US5723720A (en) 1998-03-03
JPH11507216A (ja) 1999-06-29
AU6095096A (en) 1996-12-30
ES2160250T3 (es) 2001-11-01
DK0836378T3 (da) 2001-10-01
ATE204121T1 (de) 2001-09-15
AU696959B2 (en) 1998-09-24

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